Technische Universität Dresden
Faculty of Environmental Sciences
Computational Landscape Ecology

Hülssebau, HÜL/W 432
Helmholtzstraße 10
D-01069 Dresden


Follow me on ResearchGate and Google Scholar and Computational Landscape Ecology

My primary research fields are biogeography and macroecology. I am particularly interested in evolutionary patterns and processes and the dynamics of speciation. I combine computer-based modeling techniques with genetic approaches to unravel how distributional patterns and relationships of organisms end ecological communities have been and will be shaped by climate oscillations and geological processes. My research is closely associated with invasion biology, conservation biology, and evolutionary ecology.



Research positions

Awards, grants, and scholarships

Field research experience

Professional Memberships

Invited Memberships / honors

Editorial board



Forgotten treasures – using historic collection material to assess the diversity and systematics of wide-ranging taxa

Molecular genetic approaches frequently revealed species which were considered widespread and common to harbour several narrowly distributed and potentially endangered taxa demonstrating that putatively widespread taxa need considerably more attention. However, obtaining a range-wide sampling is often unfeasible. Genetic samples obtained from scientific collections represent an appealing alternative but unfortunately age, preservation techniques, and storage conditions lead to fragmentation and contamination with exogenous DNA. Thus, historic samples were long considered useless for molecular genetic analyses. This DFG-funded project (IH133/1-1) applies specialised DNA extraction protocols, single-stranded or double-stranded library preparation, hybridization-based target enrichment, and Next Generation Sequencing (NGS) techniques suitable for short fragments to elucidate exemplarily the diversity of African Hinge-back Tortoises of the genus Kinixys from collection material (including name bearing type specimens).

Ecological modelling

Ecological models determine relationships between species, or species with their environment at various temporal and spatial scales. Depending on the research question different types of models are applied for conservation management, the management of invasive species, or to unravel patterns and processes of speciation. Niche models are frequently used to assess current habitat suitability or test refugial hypotheses and explain evolutionary processes of speciation or quantify expected future impacts of global climate and land-use change. Connectivity models assess connectivity or fragmentation of populations or suitable habitat patches for instance to evaluate the effectiveness of designated protected areas or impacts of land-use changes. Process-based dynamic models implement demographic, genetic, and dispersal data to simulate population effects e.g., to evaluate conservation measures.

Preserving the last of their kind - Conserving South African cycads

South Africa is a global hotspot for cycad diversity. Except for a single taxon all belong to the genus Encephalartos with the majority of species endemic to the country. Cycads are recognized as the most threatened plant group in the world. 69% are threatened with extinction in the near future due to poaching for ornamental purposes and traditional medicine. In close collaboration with the Cycad Specialist Group and the Wild Cycad Conservancy, this project brings together experts from botany, remote sensing, molecular genetics, conservation, and ecological modelling to conserve South Africas’ enigmatic cycads. One priority of the project is the endangered Waterberg Cycad (Encephalartos eugene-maraisii) which is endemic to the Waterberg Range located in Limpopo, South Africa, and has declined to few small and scattered subpopulations. The project combines genetics with drone surveys and remote-sensing derived modelling approaches to find additional populations, determine population connectivity, identify suitable re-introduction sites for reared seedlings, and assess the vulnerability of highly specialised Cycad-pollinator relationships to climate and land-use changes to develop appropriate conservation measures. I joined the project in 2022 and will be responsible for ecological modelling.

Ecology, Diversity, and Distribution of southern African Chelonians

The project has three main components: 1) Molecular genetics and systematics: aims at clarifying genetic relationships, identifying hidden diversity, revealing contact zones and hybridization between sympatric taxa and genetic lineages using state-of-the-art genetic techniques (mtDNA, nuclear sequences, SNPs or microsatellites). In addition, morphological data is collected to quantify shape variation using geometric morphometrics. 2) Ecological modelling: Occurrences are used to build ecological models. Depending on the research question, niche models, population connectivity models, or process-based models will be applied to identify habitat requirements, quantify niche similarity, forecast impacts of global change or simulate effects of conservation measures. 3) Ecology and natural history: Various wildlife monitoring techniques are applied. Environmental data loggers, in combination with camera traps facing burrow entrances, reveal daily and seasonal activity patterns in relation to climate. GPS and VHF tracking are used to monitor daily and seasonal movement.

Conservation & ecology

Although some chelonian species have been revealed to provide important ecosystem services, e.g. by contributing to seed dispersal and germination, the natural history of most turtles and tortoise species remains unexplored, especially in the tropics. Ecological research investigating thermal ecology, feeding ecology, and movement patterns directly contribute to successful reinforcement, reintroduction, and conservation management. My research focuses on the analyses of movement and habitat use patterns of freshwater turtles and tortoises to understand how they utilize their habitat and how they disperse. Another major aspect of my ecological research is the connectivity between populations, which, combined with knowledge on habitat use, can identify isolated populations with a higher extinction risk and guide the planning of effective conservation areas.


Phylogeographic research aims at the characterization of the spatial distribution of species or species groups and exploring the historical processes shaping it. By putting genetically distinct units that are genealogically linked through shared ancestors into a spatial context, the biogeographic history of a species or species group provides implications about historical events that have shaped the observed patterns. I apply phylogeographic techniques to detect evolutionary lineages or clusters in widespread vertebrate taxa and to subsequently explain the observed distributional patterns. In case of the Southeast Asian Snail-eating turtles of the genus Malayemys this yielded a so far unrecognized species. North American turtle species exhibit patterns of range expansion, contraction, and fragmentation during the Quaternary that are strongly congruent with phylogeographic differentiation. However, my recent research on North-American map turtles of the genus Graptemys revealed only few evolutionary lineages with little genetic diversity and therefore challenges the present inflated taxonomy of this genus.


Publications in ‘peer-reviewed’ journals

  1. Ihlow, F., Spitzweg, C., Flecks, M., Pojarkov, N., Pratyush P. Mohapatra., Fritz, U. (in press) Unexpected lack of genetic and morphological divergence in a widespread tortoise - Phylogeography of the Elongated Tortoise (Indotestudo elongata) (Blyth 1854). Salamandra,
  2. Ihlow, F., Spitzweg, C., Vamberger, M., Augustine, L., Davis, A., Leprince, B., Wagner, P., Van, T.P., Fritz, U., (2023) Molecular differentiation and conservation of the Indochinese box turtles (Cuora galbinifrons), (C. bourreti) and (C. picturata). Salamandra, 59(4), 336–346.
  3. Gallego-Garcia, N., Ihlow, F., Ettmar, S., Iverson, J.B., Fritz, U., (2023) Where to set the bar? Recent descriptions inflate species number in South American toad-headed turtles (Mesoclemmys). Zootaxa, 5263 (4), 566-574.
  4. Kehlmeier, C., Gracia, E., Ali, J.,Campbell, P., Chapman, S., Ihlow, F., Nour, E., Pierre-Huyet L., Samonds, K.E., Vences, M., Fritz, U., (2023) Ancient DNA elucidates the lost world of western Indian Ocean giant tortoises. Science Advances. 9(2), eabq2574.
  5. Bilbija, B., Spitzweg, C., Papoušek, I., Fritz, U., Földvári, G., Mullett, M., Ihlow, F.,... & Široký, P. (2022). Dermacentor reticulatus - a tick on its way from glacial refugia to a panmictic Eurasian population. International Journal for Parasitology.
  6. Fritz, U., Tolley, K., Vamberger, M., Ihlow, F. (2022). Phylogeny and phylogeography of chelonians from sub-Saharan Africa—a review of current knowledge in tribute to Margaretha D. Hofmeyr. Vertebrate Zoology. 72 951-969.
  7. Shah, A., Kayani, A., Ihlow, F., Nadeem, M., Mahmood, T., Islam, S., Päckert, M. (2022) Distribution and habitat preference of the Western Tragopan (Tragopan melanocephalus) inferred from species distribution modeling and from field data from Pakistan. Bird Conservation International. 1-14.
  8. Fritz, U., Ihlow, F. (2022). Citizen Science, taxonomy and grass snakes: iNaturalist helps to clarify variation of coloration and pattern in Natrix natrix subspecies. Vertebrate Zoology, 72, 533-549.
  9. Platt, S., Thongsavath, O., Leslie, S.C., Brakels, P., Ihlow, F., Rainwater, T. (2022) On the Occurrence of the Khorat Snail-eating Turtle (Malayemys khoratensis) in Lao, PDR with Notes on Traditional Ecological Knowledge and Exploitation. Chelonian Conservation and Biology. 21(1), 11-19.
  10. Cordero, A.G., Vamberger, M., Fritz, U., Ihlow, F. (2022) Skeletal repatterning enhances the protective capacity of the shell in African hinge-back tortoises (Kinixys). The Anatomical Record. DOI 10.1002/ar.24954.
  11. Ayaz, D., Bayrakcı, Y., Çiçek, K., Ihlow, F., Tok, C.V., Fritz, U. (2021) On the Brink of Extinction: Results of a 20-year Quest for Eiselt’s Pond Turtle (Emys orbicularis eiselti) in Southeastern Turkey. Chelonian Conservation and Biology
  12. Ihlow, F., Fritz, U., Mikulícek, P., Rödder, D. (2021) “Reconstructions of the past distribution of Testudo graeca mitochondrial lineages in the Middle East and Transcaucasia support multiple refugia since the Last Glacial Maximum”: A response to Turkozan et al. (2021). The Herpetological Journal. 31:201-203.
  13. Wagner, P., Ihlow, F., Hartmann, T., Flecks, M., Schmitz, A. & Böhme, W. (2021) Integrative approach to resolve the Calotes mystaceus Duméril & Bibron, 1837 species complex (Squamata, Agamidae). Bonn Zoological Bulletin, 70(1): 141-170.
  14. Dajcman, U., Hofmeyr, M.D., Ribeiro Anunciação, P., Ihlow, F. & Vamberger, M. (2021) Tortoise forensics: conservation genetics of the Leopard Tortoise Stigmochelys pardalis (Bell, 1828) in southern Africa. Salamandra, 57(1): 139-145.
  15. Dawson, J.E., Ihlow, F. & Platt, S.G. (2020) Malayemys subtrijuga (Schlegel and Müller 1845) - Mekong Snail-eating Turtle. In: Rhodin, A.G.J., Pritchard, P.C.H., van Dijk, P.P., Saumure, R.A., Buhlmann, K.A., Iverson, J.B., Mittermeier, R.A. (Eds). Conservation biology of freshwater turtles and tortoises: A compilation project of the IUCN/SSC tortoise and freshwater turtle specialist group. Chelonian Research Monographs, 5(14): 111.1-24.
  16. Asztalos, M., Schultze, N., Ihlow, F., Geniez, P., Berroneau, M., Delmas, C., Guiller, G., Lengentilhomme, J., Kindler, C. & Fritz, U. (2020) How often do they do it? An in-depth analysis of the hybrid zone of two European Grass Snake species (Natrix astreptophora, N. helvetica). Biological Journal of the Linnean Society, 131: 756-773.
  17. Spitzweg, C., Vamberger, M., Ihlow, F., Fritz, U. & Hofmeyr, M.D. (2020) How many species of Angulate Tortoises occur in Southern Africa? (Testudines: Testudinidae: Chersina). Zoologica Scripta, 49: 412-426.
  18. Hofmeyr, M.D., Ihlow, F., Fouche, P. & Daniels, S. (2020) Niche divergence corresponds to genetic differentiation within the Parrot-beaked Tortoise Homopus areolatus (Reptilia: Testudinidae), endemic to South Africa. Zoological Journal of the Linnean Society, 190(4): 1256-1273.
  19. Ihlow, F., Van Huyssteen, R., Vamberger, M., Cory-Toussaint, D., Hofmeyr, M.D. & Fritz, U. (2020) Geographic range extension for the Lobatse Hinge-back Tortoise Kinixys lobatsiana (Power, 1927) with first records from the Soutpansberg region. Amphibian and Reptile Conservation, 14: 132-139.
  20. Albrecht, F., Hering, J., Fuchs, E.; Illera, J.C., Ihlow, F., Shannon, T. J., Collinson, J. M., Wink, M., Martens, J. & Päckert, M. (2020) Phylogeny of the Eurasian Wren Nannus troglodytes (Aves: Passeriformes: Troglodytidae) reveals deep and complex diversification patterns of Ibero-Maghrebian and Cyrenaican populations, PlosONE: 15(3): e0230151.
  21. Vamberger, M., Ihlow, F., Asztalos, M., Dawson, J.E., Jasinski, S.E., Praschag, P. & Fritz, U. (2020) So different, yet so alike: North American slider turtles (Trachemys scripta). Vertebrate Zoology, 70(1): 87-96.
  22. Ihlow, F., Farooq, H., Qvoždík, V., Hofmeyr, M., Conradie, W., Campbell, P.D., Harvey, J., Verburgt, L. & Fritz, U. (2019). Geographic range extension of Speke‘s Hinge-back Tortoise Kinixys spekii Gray, 1863. Amphibian and Reptile Conservation, 13(2): 61-67.
  23. Geissler, P., Hartmann, T., Ihlow, F., Neang, T., Seng, R., Wagner, P., Böhme, W. (2019). Herpetofauna of the Phnom Kulen National Park, northern Cambodia — An annotated checklist. Cambodian Journal of Natural History, 40-63.
  24. Dawson, J.E., Ihlow, F., van Dijk, P.P. & K. Thirakhupt (2018) Malayemys macrocephala (Gray 1859) - Malayan Snail-eating Turtle, Rice-field Terrapin. In: Rhodin, A.G.J., Pritchard, P.C.H, van Dijk, P.P., Saumure, R.A., Buhlmann, K.A., Iverson, J.B. & R.A. Mittermeier (eds.) Conservation Biology of Freshwater Turtles and Tortoises: A Compilation Project of the IUCN/SSC Tortoise and Freshwater Turtle Specialist Group. Chelonian Research Monographs, 5(12): 108.1-16. doi:10.3854/crm.5.108.macrocephala.v1.2018
  25. Ward, M., Ihlow, F., Nadolski, B., Crane, M.S., Knierim, T., Artchawakom, T. & C.T Strine (2018) First record of male combat in the Elongated Tortoise Indotestudo elongata (Blyth, 1853) from north-eastern Thailand. Herpetology Notes, 11: 585-587. PDF
  26. Vamberger, M., Hofmeyr, M.D., Ihlow, F. & U. Fritz (2018) In quest of contact: phylogeography of helmeted terrapins (Pelomedusa galeata, P. subrufa sensu stricto). PeerJ, 6: e4901. doi:10.7717/peerj.4901
  27. Praschag, P., Ihlow, F., Flecks, M., Vamberger, M. & U. Fritz (2017) Diversity of North American map and sawback turtles (Testudines: Emydidae: Graptemys). Zoologica Scripta, 2017: 1-8. doi:10.1111/zsc.12249
  28. Courant, J., Vogt, S., Marques, R., Measey, J., Secondi, J., Rebelo, R., De Villiers, A., Ihlow, F., De Busschere, C., Backeljau, T., Rödder, D. & A. Herrel (2017) Are invasive populations characterized by a broader diet than native populations? PeerJ, 5: e3250. doi:10.7717/peerj.3250
  29. Rödder, D., Ihlow, F., Courant, J., Secondi, J, Herrel, A., Measey, G.J., Lillo, F., de Villiers, F.A., De Busschere, C. & T. Backeljau (2017) Global realized niche divergence in the African clawed frog Xenopus laevis. Ecology and Evolution, 7: 4044–4058. doi:10.1002/ece3.3010
  30. Javanbakht, H., Ihlow, F., Jablonski, D., Široký, M., Fritz, U., Rödder, D., Sharifi, M. & P. Mikulíček (2017) Genetic diversity and Quaternary range dynamics in Iranian and Transcaucasian tortoises. Biological Journal of the Linnean Society, 121: 627-640. doi:10.1093/biolinnean/blx001
  31. Vogt, S., de Villiers, F.A., Ihlow, F., Rödder, D. & J. Measey (2017) Competition and feeding ecology in two sympatric Xenopus species (Anura: Pipidae). PeerJ, 5: e3130; doi:10.7717/peerj.3130
  32. Rahn, A.K., Eßer, E., Reher, S., Ihlow, F., MacColl, A.D.C. & T.C.M. Bakker (2016) Distribution of common stickleback parasites on North Uist, Scotland, in relation to ecology and host traits. Zoology, 119: 395-402. doi:10.1016/j.zool.2016.05.009
  33. Ihlow, F., Courant, J., Secondi, J., Herrel, A., Rebelo, R., Measey, J., Lillo, F., de Villers, A., Vogt, S., Backeljau, T., de Busschere, C., Edwards, S. & D. Rödder (2016) Impacts of Climate Change on the Global Invasion Potential of the African Clawed Frog Xenopus laevis. PLoS ONE, 11(6): e0154869. doi:10.1371/journal.pone.0154869
  34. Ihlow, F., Dawson, J.E., Hartmann, T. & S. Som (2016) Indotestudo elongata (Blyth, 1854) – Elongated Tortoise. In: Rhodin, A.G.J., Pritchard, P.C.H, van Dijk, P.P., Saumure, R.A., Buhlmann, K.A., Iverson, J.B. & R.A. Mittermeier (eds.) Conservation Biology of Freshwater Turtles and Tortoises: A Compilation Project of the IUCN/SSC Tortoise and Freshwater Turtle Specialist Group. Chelonian Research Monographs, 5(9): 096.1–14. doi:10.3854/crm.5.096.elongata.v1.2016
  35. Ihlow, F., Vamberger, M., Flecks, M., Hartmann, T., Cota, M., Makchai, S., Meewattana, P., Dawson, J.E., Kheng, L., Rödder, D. & U. Fritz (2016) Integrative Taxonomy of Southeast Asian Snail-Eating Turtles (Geoemydidae: Malayemys) Reveals a New Species and Mitochondrial Introgression. PLoS ONE, 11(4): e0153108. doi:10.1371/journal.pone.0153108
  36. Ahmadzadeh, F., Flecks, M., Carretero, M.A., Böhme, W., Ihlow F., Kapli, P., Miraldo, A. & D. Rödder (2016) Separate histories in both sides of the Mediterranean: Phylogeny and niche evolution of ocellated lizards. Journal of Biogeography, 43: 1242–1253. doi:10.1111/jbi.12703
  37. Slavenko, A., Itescu, Y., Ihlow, F. & S. Meiri (2016) Home is where the shell is: predicting turtle home range sizes. Journal of Animal Ecology, 85: 106-114. doi:10.1111/1365-2656.12446
  38. Geissler, P., Hartmann, T., Ihlow, F., Rödder, D., Poyarkov, N.A. Jr., Nguyen, T.Q., Ziegler, T. & W. Böhme (2015) The Lower Mekong: an insurmountable barrier to amphibians in southern Indochina? Biological Journal of the Linnean Society. doi:10.1111/bij.12444
  39. Ghaffari, H., Ihlow, F., Plummer, M.V., Karami, M., Khorasani, N., Mahroo, B.S. & D. Rödder (2014) Home range and habitat selection of the endangered Euphrates Softshell Turtle Rafetus euphraticus (Daudin, 1802) in a fragmented habitat in southwestern Iran. Chelonian Conservation and Biology, 13: 202-215. doi:10.2744/CCB-1071.1
  40. Hartmann, T., Betts, A.B., De Greef, S. & F. Ihlow (2014) First record of the rare parachute gecko Ptychozoon trinotaterra Brown, 1999 from Cambodia. Cambodian Journal of Natural History, 09/2014: 12-13.
  41. Ihlow, F., Rödder, D., Bochynek, T., Sovath, S., Handschuh, M. & W. Böhme (2014) Reinforcement as a conservation tool - assessing site fidelity and movement of the endangered elongated tortoise Indotestudo elongata (Blyth, 1854). Journal of Natural History, 48: 2473-2485. doi:10.1080/00222933.2014.925595
  42. Ihlow, F., Bonke, R., Hartmann, T., Geissler, P., Behler, N. & D. Rödder (2014) Habitat suitability, coverage by protected areas and population connectivity for the Siamese crocodile Crocodylus siamensis Schneider, 1801. Aquatic Conservation, 25: 544-554. doi:10.1002/aqc.2473
  43. Ihlow, F., Ahmadzadeh, F., Ghaffari, H., Taskavak, E., Hartmann, T., Etzbauer, C. & D. Rödder (2014) Assessment of genetic structure, habitat suitability and effectiveness of reserves for future conservation planning of the Euphrates Soft-shelled Turtle Rafetus euphraticus (Daudin, 1802). Aquatic Conservation, 24: 831-840. doi:10.1002/aqc.2454
  44. van Schingen, M., Ihlow, F., Nguyen, T.Q., Ziegler, T., Bonkowski, M., Wu, Z. & D. Rödder (2014) Potential distribution and effectiveness of the protected area network for the crocodile lizard, Shinisaurus crocodilurus (Reptilia: Squamata: Sauria). Salamandra, 50: 71-76. PDF
  45. Bonke, R., Ihlow, F., Böhme, W. & D. Rödder (2014) Movement patterns of Tomistoma schlegelii in the Sekonyer Kanan River (Tanjung Puting National Park, Central Kalimantan, Indonesia): preliminary range size estimates. Salamandra, 50: 40-52. PDF
  46. Rödder, D. & Ihlow, F. (2013) Chelonians in a changing climate: can nest site selection prevent sex ratio skews? Animal Conservation, 16: 491-492. doi:10.1111/acv.12079
  47. Rödder, D., Lawing, A.M., Flecks, M., Ahmadzadeh, F., Dambach, J., Engler, J.O., Habel, J.C., Hartmann, T., Hörnes, D., Ihlow, F., Schidelko, K., Stiels, D. & Polly, P.D. (2013) Evaluating the significance of paleophylogeographic species distribution models in reconstructing quaternary range-shifts of Nearctic chelonians. PLoS ONE, 8(10): e72855. doi:10.1371/journal.pone.0072855
  48. Hartmann, T., Ihlow, F., Edwards, S., Sovath, S., Handschuh, M. & W. Böhme (2013) A Preliminary Annotated Checklist of the Amphibians and Reptiles of the Kulen Promtep Wildlife Sanctuary in Northern Cambodia. Asian Herpetological Research, 4: 36-55.
  49. Hartmann, T., Geissler, P., Poyarkov, N.A., Ihlow, F., Galoyan, E.A., Rödder, D. & W. Böhme (2013) A new species of the genus Calotes Cuvier, 1817 (Squamata: Agamidae) from southern Vietnam. Zootaxa, 3599: 246-260. doi:10.11646/zootaxa.3599.3.3
  50. Ihlow, F., Dambach, J., Engler, J., Flecks, M., Hartmann, T., Nekum, S., Rajaei, H. & D. Rödder (2012) On the brink of extinction? How climate change may affect global Chelonian species richness and distribution. Global Change Biology, 18: 1520-1530. doi:10.1111/j.1365-2486.2011.02623.x
  51. Ihlow, F., Geissler P., Sovath S., Handschuh, M. & W. Böhme (2012) Feeding ecology of Indotestudo elongata (Blyth 1853) in the wild in Cambodia and Vietnam. Herpetology Notes, 5: 5-7. PDF

Selected publications in other journals

  1. Ihlow, F. & J. Dawson (2016) Ecological research and conservation status of turtles at the Tonlé Sap Biosphere Reserve in central Cambodia. Testudo, 8: 13-25.
  2. Ihlow, F. (2016) 1, 2, oder 3 - Diversität und Verbreitung der Sumpfschildkröten der Gattung Malayemys. Sacalia, 14: 39-50.
  3. Ihlow, F. & J. Dawson (2016) Fang und Handel der aquatischen Schildkröten im Tonlé Sap Biosphären Reservat in Zentralkambodscha. Schildkröten im Fokus, 13: 23-31.
  4. Ihlow, F. (2015) Bewegingspatronen en voedingsecologie van de Maleisische slakkeneter (Malayemys subtrijuga) Schlegel & Müller, 1844. Trionyx, 12: 118-124.
  5. Ihlow, F. (2013) Geographische Variation der Gelbkopf-Landschildkröte Indotestudo elongata (Blyth, 1854). Testudo, 22: 7-19.
  6. Ihlow, F. (2011) Die Schildkrötenfauna Kambodschas. Schildkröten im Fokus, 8: 23-35.
  7. Ihlow, F. (2011) Kambodscha, ein Königreich der Wunder - Feldherpetologische Arbeit an Indotestudo elongata. Marginata, 8: 29-41.
  8. Ihlow, F. & M. Handschuh (2011) Auswilderung von Indotestudo elongata im Kulen Promtep Wildlife Sanctuary im Norden Kambodschas. Marginata, 8(4): 16-23.
  9. Ihlow, F., Handschuh, M. & M. Raffel (2011) Haltung und Nachzucht der Gelbkopf-Landschildkröte (Indotestudo elongata) im Angkor Centre for Conservation of Biodiversity (ACCB), Kambodscha. Marginata, 8(4): 24-30.